Estimating Abundance From One-Dimensional Passive Acoustic Surveys

ABSTRACT Conventional distance sampling, the most-used method of estimating animal density and abundance, requires ranges to detected individuals, which are not easily measured for vocalizations. However, in some circumstances the sequential pattern of detection of vocalizations along a transect line gives information about the range of detection. Thus, from a one-dimensional acoustic point-transect survey (i.e., records of vocalizations detected or not detected at regularly spaced listening stations) it is possible to obtain a useful estimate of density or abundance. I developed equations for estimation of density for one-dimensional surveys. Using simulations I found that for the method to have little bias when both range of detection and rate of vocalization need to be estimated, stations needed to be spaced at 30–80% of the range of detection and the rate of vocalization should be >0.7. If either the range of detection or rate of vocalization is known, conditions are relaxed, and when both parameters are known the method works well almost universally. In favorable conditions for one-dimensional methods, estimated abundances have overall errors not much larger than those from conventional line-transect distance sampling. The methods appeared useful when applied to real acoustic data from whale surveys. The techniques may also be useful in surveys with nonacoustic detection of animals.     

Evaluation of three methods to estimate density and detectability from roadside point counts

Roadside point counts are often used to estimate trends of bird populations. The use of aural counts of birds without adjustment for detection probability, however, can lead to incorrect population trend estimates. We compared precision of estimates of density and detectability of whistling northern bobwhites (Colinus virginianus) using distance sampling, independent double-observer, and removal methods from roadside surveys. Two observers independently recorded each whistling bird heard, distance from the observer, and time of first detection at 362 call-count stops in Ohio. We examined models that included covariates for year and observer effects for each method and distance from observer effects for the double-observer and removal methods using Akaike's Information Criterion (AIC). The best model of detectability from distance sampling included observer and year effects. The best models from the removal and double-observer techniques included observer and distance effects. All 3 methods provided precise estimates of detection probability (CV = 2.4–4.4%) with a range of detectability of 0.44–0.95 for a 6-min survey. Density estimates from double-observer surveys had the lowest coefficient of variation (2005 = 3.2%, 2006 = 1.7%), but the removal method also provided precise estimates of density (2005 CV = 3.4%, 2006 CV = 4.8%), and density estimates from distance sampling were less precise (2005 CV = 9.6%, 2006 CV = 7.9%). Assumptions of distance sampling were violated in our study because probability of detecting bobwhites near the observer was <1 or the roadside survey points were not randomly distributed with respect to the birds. Distances also were not consistently recorded by individual members of observer pairs. Although double-observer surveys provided more precise estimates, we recommend using the removal method to estimate detectability and abundance of bobwhites. The removal method provided precise estimates of density and detection probability and requires half the personnel time as double-observer surveys. Furthermore, the likelihood of meeting model assumptions is higher for the removal survey than with independent double-observers. © 2011 The Wildlife Society.     

Using soundscape recordings to estimate bird species abundance, richness, and composition

ABSTRACT Point counts are the most frequently used technique for sampling bird populations and communities, but have well‐known limitations such as inter‐ and intraobserver errors and limited availability of expert field observers. The use of acoustic recordings to survey birds offers solutions to these limitations. We designed a Soundscape Recording System (SRS) that combines a four‐channel, discrete microphone system with a quadraphonic playback system for surveying bird communities. We compared the effectiveness of SRS and point counts for estimating species abundance, richness, and composition of riparian breeding birds in California by comparing data collected simultaneously using both methods. We used the temporal‐removal method to estimate individual bird detection probabilities and species abundances using the program MARK. Akaike's Information Criterion provided strong evidence that detection probabilities differed between the two survey methods and among the 10 most common species. The probability of detecting birds was higher when listening to SRS recordings in the laboratory than during the field survey. Additionally, SRS data demonstrated a better fit to the temporal‐removal model assumptions and yielded more reliable estimates of detection probability and abundance than point‐count data. Our results demonstrate how the perceptual constraints of observers can affect temporal detection patterns during point counts and thus influence abundance estimates derived from time‐of‐detection approaches. We used a closed‐population capture–recapture approach to calculate jackknife estimates of species richness and average species detection probabilities for SRS and point counts using the program CAPTURE. SRS and point counts had similar species richness and detection probabilities. However, the methods differed in the composition of species detected based on Jaccard's similarity index. Most individuals (83%) detected during point counts vocalized at least once during the survey period and were available for detection using a purely acoustic technique, such as SRS. SRS provides an effective method for surveying bird communities, particularly when most species are detected by sound. SRS can eliminate or minimize observer biases, produce permanent records of surveys, and resolve problems associated with the limited availability of expert field observers.     

Red-Shouldered Hawk Occupancy Surveys in Central Minnesota,USA

ABSTRACT Forest-dwelling raptors are often difficult to detect because many species occur at low density or are secretive. Broadcasting conspecific vocalizations can increase the probability of detecting forest-dwelling raptors and has been shown to be an effective method for locating raptors and assessing their relative abundance. Recent advances in statistical techniques based on presence—absence data use probabilistic arguments to derive probability of detection when it is < 1 and to provide a model and likelihood-based method for estimating proportion of sites occupied. We used these maximum-likelihood models with data from red-shouldered hawk (Buteo lineatus) call-broadcast surveys conducted in central Minnesota, USA, in 1994–1995 and 2004–2005. Our objectives were to obtain estimates of occupancy and detection probability 1) over multiple sampling seasons (yr), 2) incorporating within-season time-specific detection probabilities, 3) with call type and breeding stage included as covariates in models of probability of detection, and 4) with different sampling strategies. We visited individual survey locations 2–9 times per year, and estimates of both probability of detection (range = 0.28-0.54) and site occupancy (range = 0.81-0.97) varied among years. Detection probability was affected by inclusion of a within-season time-specific covariate, call type, and breeding stage. In 2004 and 2005 we used survey results to assess the effect that number of sample locations, double sampling, and discontinued sampling had on parameter estimates. We found that estimates of probability of detection and proportion of sites occupied were similar across different sampling strategies, and we suggest ways to reduce sampling effort in a monitoring program.     

Comparison of methods for estimating density of forest songbirds from point counts

New analytical methods have been promoted for estimating the probability of detection and density of birds from count data but few studies have compared these methods using real data. We compared estimates of detection probability and density from distance and time-removal models and survey protocols based on 5- or 10-min counts and outer radii of 50 or 100 m. We surveyed singing male Acadian flycatchers (Empidonax virescens), cerulean warblers (Dendroica cerulea), Kentucky warblers (Oporornis formosus), Louisiana waterthrushes (Parkesia motacilla), wood thrushes (Hylocichla mustelina), and worm-eating warblers (Helmitheros vermivorum) in bottomland and upland forest across 5 states in the Central Hardwoods Bird Conservation Region during the breeding season in 2007 and 2008. Detection probabilities differed between distance and time-removal models and species detectabilities were affected differently by year, forest type, and state. Density estimates from distance models were generally higher than from time-removal models, resulting from lower detection probabilities estimated by distance models. We found support for individual heterogeneity (modeled as a finite mixture model) in the time-removal models and that 50-m radius counts generated density estimates approximately twice as high as 100-m radius counts. Users should be aware that in addition to estimating different components of detectability, density estimates derived from distance and time-removal models can be affected by survey protocol because some count durations and plot radii may better meet model assumptions than others. The choice of a method may not affect the use of estimates for relative comparisons (e.g., when comparing habitats) but could affect conclusions when used to estimate population size. We recommend careful consideration of assumptions when deciding on point-count protocol and selection of analysis methods. © 2011 The Wildlife Society.     

The Optimal Number of Surveys when Detectability Varies

The survey of plant and animal populations is central to undertaking field ecology. However, detection is imperfect, so the absence of a species cannot be determined with certainty. Methods developed to account for imperfect detectability during surveys do not yet account for stochastic variation in detectability over time or space. When each survey entails a fixed cost that is not spent searching (e.g., time required to travel to the site), stochastic detection rates result in a trade-off between the number of surveys and the length of each survey when surveying a single site. We present a model that addresses this trade-off and use it to determine the number of surveys that: 1) maximizes the expected probability of detection over the entire survey period; and 2) is most likely to achieve a minimally-acceptable probability of detection. We illustrate the applicability of our approach using three practical examples (minimum survey effort protocols, number of frog surveys per season, and number of quadrats per site to detect a plant species) and test our model''s predictions using data from experimental plant surveys. We find that when maximizing the expected probability of detection, the optimal survey design is most sensitive to the coefficient of variation in the rate of detection and the ratio of the search budget to the travel cost. When maximizing the likelihood of achieving a particular probability of detection, the optimal survey design is most sensitive to the required probability of detection, the expected number of detections if the budget were spent only on searching, and the expected number of detections that are missed due to travel costs. We find that accounting for stochasticity in detection rates is likely to be particularly important for designing surveys when detection rates are low. Our model provides a framework to do this.     

Site occupancy models with heterogeneous detection probabilities

Models for estimating the probability of occurrence of a species in the presence of imperfect detection are important in many ecological disciplines. In these "site occupancy" models, the possibility of heterogeneity in detection probabilities among sites must be considered because variation in abundance (and other factors) among sampled sites induces variation in detection probability (p). In this article, I develop occurrence probability models that allow for heterogeneous detection probabilities by considering several common classes of mixture distributions for p. For any mixing distribution, the likelihood has the general form of a zero-inflated binomial mixture for which inference based upon integrated likelihood is straightforward. A recent paper by Link demonstrates that in closed population models used for estimating population size, different classes of mixture distributions are indistinguishable from data, yet can produce very different inferences about population size. I demonstrate that this problem can also arise in models for estimating site occupancy in the presence of heterogeneous detection probabilities. The implications of this are discussed in the context of an application to avian survey data and the development of animal monitoring programs.     

Determination of contact angle by molecular simulation using number and atomic density contours

The contact angles of Lennard-Jones fluid droplets on a structureless solid surface, simulated using Monte Carlo simulation, are calculated by fitting isochoric surfaces and making a number of assumptions about the droplet. The results show that there are significant uncertainties in the calculated contact angles due to the choice of these assumptions, such as the grid size used in tracking the isochoric density profile, the omission of isochoric data points near the surface and the function used to fit the isochoric profile. In this study, we propose a new method of calculating density contours based on atomic density instead of number density. This method results in a much smaller variation in contact angle when applying different assumptions than using number density for isochoric contours. The most consistent results, across a range of assumptions about the droplet and the contact angle, come from averaging the contact angle from several isochoric density profiles. In addition, this gives a measurement of the variation due to the choice of isochoric density.     

Dealing with detection error in site occupancy surveys: what can we do with a single survey?

Aim Site occupancy probabilities of target species are commonly used in various ecological studies, e.g. to monitor current status and trends in biodiversity. Detection error introduces bias in the estimators of site occupancy. Existing methods for estimating occupancy probability in the presence of detection error use replicate surveys. These methods assume population closure, i.e. the site occupancy status remains constant across surveys, and independence between surveys. We present an approach for estimating site occupancy probability in the presence of detection error that requires only a single survey and does not require assumption of population closure or independence. In place of the closure assumption, this method requires covariates that affect detection and occupancy.Methods Penalized maximum-likelihood method was used to estimate the parameters. Estimability of the parameters was checked using data cloning. Parametric boostrapping method was used for computing confidence intervals.Important findings The single-survey approach facilitates analysis of historical datasets where replicate surveys are unavailable, situations where replicate surveys are expensive to conduct and when the assumptions of closure or independence are not met. This method saves significant amounts of time, energy and money in ecological surveys without sacrificing statistical validity. Further, we show that occupancy and habitat suitability are not synonymous and suggest a method to estimate habitat suitability using single-survey data.     

A spatially explicit capture–recapture estimator for single‐catch traps

Single‐catch traps are frequently used in live‐trapping studies of small mammals. Thus far, a likelihood for single‐catch traps has proven elusive and usually the likelihood for multicatch traps is used for spatially explicit capture–recapture (SECR) analyses of such data. Previous work found the multicatch likelihood to provide a robust estimator of average density. We build on a recently developed continuous‐time model for SECR to derive a likelihood for single‐catch traps. We use this to develop an estimator based on observed capture times and compare its performance by simulation to that of the multicatch estimator for various scenarios with nonconstant density surfaces. While the multicatch estimator is found to be a surprisingly robust estimator of average density, its performance deteriorates with high trap saturation and increasing density gradients. Moreover, it is found to be a poor estimator of the height of the detection function. By contrast, the single‐catch estimators of density, distribution, and detection function parameters are found to be unbiased or nearly unbiased in all scenarios considered. This gain comes at the cost of higher variance. If there is no interest in interpreting the detection function parameters themselves, and if density is expected to be fairly constant over the survey region, then the multicatch estimator performs well with single‐catch traps. However if accurate estimation of the detection function is of interest, or if density is expected to vary substantially in space, then there is merit in using the single‐catch estimator when trap saturation is above about 60%. The estimator's performance is improved if care is taken to place traps so as to span the range of variables that affect animal distribution. As a single‐catch likelihood with unknown capture times remains intractable for now, researchers using single‐catch traps should aim to incorporate timing devices with their traps.     

Inferring species richness using multispecies occupancy modeling: Estimation performance and interpretation

Multispecies occupancy models can estimate species richness from spatially replicated multispecies detection/non‐detection survey data, while accounting for imperfect detection. A model extension using data augmentation allows inferring the total number of species in the community, including those completely missed by sampling (i.e., not detected in any survey, at any site). Here we investigate the robustness of these estimates. We review key model assumptions and test performance via simulations, under a range of scenarios of species characteristics and sampling regimes, exploring sensitivity to the Bayesian priors used for model fitting. We run tests when assumptions are perfectly met and when violated. We apply the model to a real dataset and contrast estimates obtained with and without predictors, and for different subsets of data. We find that, even with model assumptions perfectly met, estimation of the total number of species can be poor in scenarios where many species are missed (>15%–20%) and that commonly used priors can accentuate overestimation. Our tests show that estimation can often be robust to violations of assumptions about the statistical distributions describing variation of occupancy and detectability among species, but lower‐tail deviations can result in large biases. We obtain substantially different estimates from alternative analyses of our real dataset, with results suggesting that missing relevant predictors in the model can result in richness underestimation. In summary, estimates of total richness are sensitive to model structure and often uncertain. Appropriate selection of priors, testing of assumptions, and model refinement are all important to enhance estimator performance. Yet, these do not guarantee accurate estimation, particularly when many species remain undetected. While statistical models can provide useful insights, expectations about accuracy in this challenging prediction task should be realistic. Where knowledge about species numbers is considered truly critical for management or policy, survey effort should ideally be such that the chances of missing species altogether are low.     

Effects of branch length uncertainty on Bayesian posterior probabilities for phylogenetic hypotheses

In Bayesian phylogenetics, confidence in evolutionary relationships is expressed as posterior probability--the probability that a tree or clade is true given the data, evolutionary model, and prior assumptions about model parameters. Model parameters, such as branch lengths, are never known in advance; Bayesian methods incorporate this uncertainty by integrating over a range of plausible values given an assumed prior probability distribution for each parameter. Little is known about the effects of integrating over branch length uncertainty on posterior probabilities when different priors are assumed. Here, we show that integrating over uncertainty using a wide range of typical prior assumptions strongly affects posterior probabilities, causing them to deviate from those that would be inferred if branch lengths were known in advance; only when there is no uncertainty to integrate over does the average posterior probability of a group of trees accurately predict the proportion of correct trees in the group. The pattern of branch lengths on the true tree determines whether integrating over uncertainty pushes posterior probabilities upward or downward. The magnitude of the effect depends on the specific prior distributions used and the length of the sequences analyzed. Under realistic conditions, however, even extraordinarily long sequences are not enough to prevent frequent inference of incorrect clades with strong support. We found that across a range of conditions, diffuse priors--either flat or exponential distributions with moderate to large means--provide more reliable inferences than small-mean exponential priors. An empirical Bayes approach that fixes branch lengths at their maximum likelihood estimates yields posterior probabilities that more closely match those that would be inferred if the true branch lengths were known in advance and reduces the rate of strongly supported false inferences compared with fully Bayesian integration.     

Testing the consistency of wildlife data types before combining them: the case of camera traps and telemetry

Wildlife data gathered by different monitoring techniques are often combined to estimate animal density. However, methods to check whether different types of data provide consistent information (i.e., can information from one data type be used to predict responses in the other?) before combining them are lacking. We used generalized linear models and generalized linear mixed-effects models to relate camera trap probabilities for marked animals to independent space use from telemetry relocations using 2 years of data for fishers (Pekania pennanti) as a case study. We evaluated (1) camera trap efficacy by estimating how camera detection probabilities are related to nearby telemetry relocations and (2) whether home range utilization density estimated from telemetry data adequately predicts camera detection probabilities, which would indicate consistency of the two data types. The number of telemetry relocations within 250 and 500 m from camera traps predicted detection probability well. For the same number of relocations, females were more likely to be detected during the first year. During the second year, all fishers were more likely to be detected during the fall/winter season. Models predicting camera detection probability and photo counts solely from telemetry utilization density had the best or nearly best Akaike Information Criterion (AIC), suggesting that telemetry and camera traps provide consistent information on space use. Given the same utilization density, males were more likely to be photo-captured due to larger home ranges and higher movement rates. Although methods that combine data types (spatially explicit capture–recapture) make simple assumptions about home range shapes, it is reasonable to conclude that in our case, camera trap data do reflect space use in a manner consistent with telemetry data. However, differences between the 2 years of data suggest that camera efficacy is not fully consistent across ecological conditions and make the case for integrating other sources of space-use data.     

Designing occupancy surveys and interpreting non‐detection when observations are imperfect

Aim Conservation practitioners use biological surveys to ascertain whether or not a site is occupied by a particular species. Widely used statistical methods estimate the probability that a species will be detected in a survey of an occupied site. However, these estimates of detection probability are alone not sufficient to calculate the probability that a species is present given that it was not detected. The aim of this paper is to demonstrate methods for correctly calculating (1) the probability a species occupies a site given one or more non‐detections, and (2) the number of sequential non‐detections necessary to assert, with a pre‐specified confidence, that a species is absent from a site. Location Occupancy data for a tree frog in eastern Australia serve to illustrate methods that may be applied anywhere species’ occupancy data are used and detection probabilities are < 1. Methods Building on Bayesian expressions for the probability that a site is occupied by a species when it is not detected, and the number of non‐detections necessary to assert absence with a pre‐specified confidence, we estimate occupancy probabilities across tree frog survey locations, drawing on information about where and when the species was detected during surveys. Results We show that the number of sequential non‐detections necessary to assert that a species is absent increases nonlinearly with the prior probability of occupancy, the probability of detection if present, and the desired level of confidence about absence. Main conclusions If used more widely, the Bayesian analytical approaches illustrated here would improve collection and interpretation of biological survey data, providing a coherent way to incorporate detection probability estimates in the design of minimum survey requirements for monitoring, impact assessment and distribution modelling.     

On the importance of g(0) for estimating bird population densities with standard distance-sampling: implications from a telemetry study and a literature review

Raw count data are often used to estimate bird population densities. However, such data do not consider detection probability. As an alternative, methods that model detection probability such as distance-sampling have been proposed. However, standard distance-sampling provides reliable estimates for absolute density only when the underlying assumptions are met. One of the most critical of these assumptions is that animals on a transect line or at an observation point have to be detected with certainty (the g (0) = 1 assumption). We radiotagged nine Orphean Warblers Sylvia hortensis and estimated their short-distance detection probability. Birds were radio-located in 264 cases in single bushes or trees. Their visual detection probability after a 5-min search was only 0.58 (sd = ±0.14, range = 0.38–0.80), although the observer knew the bird's location. Furthermore, we carried out a literature review to assess how the g (0) = 1 assumption is handled in practice. None of the 28 standard distance-sampling papers reviewed contained an estimation of g (0). In 57% of the papers, the g (0) = 1 assumption was not even mentioned. Nevertheless, none of the authors declared their estimates as being relative. Our empirical data show that the g (0) = 1 assumption would be severely violated for a foliage-gleaning bird species at a desert stopover site outside the breeding season. The literature review revealed that the testing of the g (0) = 1 assumption is largely ignored in practice. We strongly suggest that more attention should be paid to the testing of this key assumption, because results may not be reliable when it is violated. If it is not possible to test the g (0) = 1 assumption or g (0) is less than 1, alternative methods should be used. Another possibility is to estimate detection probability by the means of radiotagged individuals.     

Optimising camera trap surveys for the Carpentarian Pseudantechinus (Pseudantechinus mimulus) in northwest Queensland

The Carpentarian Pseudantechinus (Pseudantechinus mimulus) is a poorly studied dasyurid marsupial that inhabits rocky outcrops in the Mount Isa Inlier bioregion in Queensland and the Gulf Coastal and Gulf Fall and Uplands bioregions in the Northern Territory. It is readily detected by passive infrared triggered camera traps (‘camera traps’). Camera trap data can be used to develop detection probability estimates from which activity patterns can be inferred, but no effort has previously been made to determine changes in the detectability of P. mimulus throughout the year. We undertook a 13-month baited camera trap survey across nine sampling periods at 60 locations of known historic presence or nearby suitable habitat to assess the change in detection rates and detection probabilities of P. mimulus across a year. Detection probabilities were calculated from camera trap data within a single-species, multi-season occupancy framework to determine optimal survey timing. Detection probability data were used to calculate the likelihood of false absences to determine optimal survey duration. We recorded 2493 detections of P. mimulus over 10 966 camera days. Detection probability ranged from 0.009 to 0.179 and was significantly higher from April to October than from November to March. The likelihood of false absences varied by sampling period and desired level of confidence. We find that camera trap surveys for P. mimulus are best conducted from April to October, but optimal survey duration is dependent upon the time of year and desired level of confidence that an observed absence from a given site reflects a true absence at that site. Attaining a minimum of 80% confidence of absence requires as few as 9 days of survey effort in May to 16 days of survey effort in October.     

Order‐Restricted Semiparametric Inference for the Power Bias Model

Summary The power bias model, a generalization of length‐biased sampling, is introduced and investigated in detail. In particular, attention is focused on order‐restricted inference. We show that the power bias model is an example of the density ratio model, or in other words, it is a semiparametric model that is specified by assuming that the ratio of several unknown probability density functions has a parametric form. Estimation and testing procedures under constraints are developed in detail. It is shown that the power bias model can be used for testing for, or against, the likelihood ratio ordering among multiple populations without resorting to any parametric assumptions. Examples and real data analysis demonstrate the usefulness of this approach.     

Can you hear me now? Range‐testing a submerged passive acoustic receiver array in a Caribbean coral reef habitat

Submerged passive acoustic technology allows researchers to investigate spatial and temporal movement patterns of many marine and freshwater species. The technology uses receivers to detect and record acoustic transmissions emitted from tags attached to an individual. Acoustic signal strength naturally attenuates over distance, but numerous environmental variables also affect the probability a tag is detected. Knowledge of receiver range is crucial for designing acoustic arrays and analyzing telemetry data. Here, we present a method for testing a relatively large‐scale receiver array in a dynamic Caribbean coastal environment intended for long‐term monitoring of multiple species. The U.S. Geological Survey and several academic institutions in collaboration with resource management at Buck Island Reef National Monument (BIRNM), off the coast of St. Croix, recently deployed a 52 passive acoustic receiver array. We targeted 19 array‐representative receivers for range‐testing by submersing fixed delay interval range‐testing tags at various distance intervals in each cardinal direction from a receiver for a minimum of an hour. Using a generalized linear mixed model (GLMM), we estimated the probability of detection across the array and assessed the effect of water depth, habitat, wind, temperature, and time of day on the probability of detection. The predicted probability of detection across the entire array at 100 m distance from a receiver was 58.2% (95% CI: 44.0–73.0%) and dropped to 26.0% (95% CI: 11.4–39.3%) 200 m from a receiver indicating a somewhat constrained effective detection range. Detection probability varied across habitat classes with the greatest effective detection range occurring in homogenous sand substrate and the smallest in high rugosity reef. Predicted probability of detection across BIRNM highlights potential gaps in coverage using the current array as well as limitations of passive acoustic technology within a complex coral reef environment.     

Using density surface models to estimate spatio-temporal changes in population densities and trend

Precise measures of population abundance and trend are needed for species conservation; these are most difficult to obtain for rare and rapidly changing populations. We compare uncertainty in densities estimated from spatio–temporal models with that from standard design-based methods. Spatio–temporal models allow us to target priority areas where, and at times when, a population may most benefit. Generalised additive models were fitted to a 31-year time series of point-transect surveys of an endangered Hawaiian forest bird, the Hawai‘i ‘ākepa Loxops coccineus. This allowed us to estimate bird densities over space and time. We used two methods to quantify uncertainty in density estimates from the spatio–temporal model: the delta method (which assumes independence between detection and distribution parameters) and a variance propagation method. With the delta method we observed a 52% decrease in the width of the design-based 95% confidence interval (CI), while we observed a 37% decrease in CI width when propagating the variance. We mapped bird densities as they changed across space and time, allowing managers to evaluate management actions. Integrating detection function modelling with spatio–temporal modelling exploits survey data more efficiently by producing finer-grained abundance estimates than are possible with design-based methods as well as producing more precise abundance estimates. Model-based approaches require switching from making assumptions about the survey design to assumptions about bird distribution. Such a switch warrants consideration. In this case the model-based approach benefits conservation planning through improved management efficiency and reduced costs by taking into account both spatial shifts and temporal changes in population abundance and distribution.     

Evaluating Methods for Counting Cryptic Carnivores

ABSTRACT Numerous techniques have been proposed to estimate carnivore abundance and density, but few have been validated against populations of known size. We used a density estimate established by intensive monitoring of a population of radiotagged leopards (Panthera pardus) with a detection probability of 1.0 to evaluate efficacy of track counts and camera-trap surveys as population estimators. We calculated densities from track counts using 2 methods and compared performance of 10 methods for calculating the effectively sampled area for camera-trapping data. Compared to our reference density (7.33 ± 0.44 leopards/100 km²), camera-trapping generally produced more accurate but less precise estimates than did track counts. The most accurate result (6.97 ± 1.88 leopards/100 km²) came from camera-trap data with a sampled area buffered by a boundary strip representing the mean maximum distance moved by leopards outside the survey area (MMDMOSA) established by telemetry. However, contrary to recent suggestions, the traditional method of using half the mean maximum distance moved from photographic recaptures did not result in gross overestimates of population density (6.56 ± 1.92 leopards/100 km²) but rather displayed the next best performance after MMDMOSA. The only track-count method comparable to reference density employed a capture-recapture framework applied to data when individuals were identified from their tracks (6.45 ± 1.43 leopards/100 km²) but the underlying assumptions of this technique limit more widespread application. Our results demonstrate that if applied correctly, camera-trap surveys represent the best balance of rigor and cost-effectiveness for estimating abundance and density of cryptic carnivore species that can be identified individually.